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 THE ROUTLEDGE HANDBOOK
OF ARABIC SECOND LANGUAGE
ACQUISITION

The Routledge Handbook of Arabic Second Language Acquisition introduces major current
approaches in Arabic second language acquisition (SLA) research and offers empirical findings
on crucial aspects and issues to do with the learning of Arabic as a foreign language and Arabic
SLA. It brings together leading academics in the field to synthesize existing research and
develops a new framework for analyzing important topics within Arabic SLA.

This handbook will be suitable as a reference work for advanced undergraduate and
postgraduate students and scholars actively researching in this area and is primarily relevant to
sister disciplines within teacher training and Arabic applied linguistics. The themes and
findings should, however, also be attractive to other areas of study, including theoretical
linguistics, psycholinguistics, cognition, and cognitive psychology.

Mohammad T. Alhawary is Professor of Arabic linguistics and second language acquisition

at the University of Michigan, where he teaches courses on both Arabic language and Arabic
theoretical and applied linguistics.
 THE ROUTLEDGE
HANDBOOK OF ARABIC
SECOND LANGUAGE
ACQUISITION

Edited by
Mohammad T. Alhawary
 First published 2018
by Routledge
2 Park Square, Milton Park, Abingdon, Oxon OX14 4RN
and by Routledge
711 Third Avenue, New York, NY 10017
Routledge is an imprint of the Taylor & Francis Group, an informa business
© 2018 selection and editorial matter, Mohammad T. Alhawary; individual chapters, the contributors
The right of Mohammad T. Alhawary to be identified as the author of the editorial material,
and of the authors for their individual chapters, has been asserted in accordance with sections 77
and 78 of the Copyright, Designs and Patents Act 1988.
All rights reserved. No part of this book may be reprinted or reproduced or utilised in any
form or by any electronic, mechanical, or other means, now known or hereafter invented,
including photocopying and recording, or in any information storage or retrieval system,
without permission in writing from the publishers.
Trademark notice: Product or corporate names may be trademarks or registered trademarks,
and are used only for identification and explanation without intent to infringe.
British Library Cataloguing-in-Publication Data
A catalogue record for this book is available from the British Library
Library of Congress Cataloging-in-Publication Data
A catalog record for this book has been requested
ISBN: 978-1-138-94055-0 (hbk)
ISBN: 978-1-315-67426-1(ebk)
Typeset in Times New Roman
by Apex CoVantage, LLC
 CONTENTS

List of contributors viii

Acknowledgmentsx

Introduction1

PART I
Arabic L2 phonology and phonetics 7

1 Frequency and L1 transfer effects for the perception and production

of Arabic lexical stress by L1 English and L1 Chinese learners of
Arabic as an L2 9
Cheng-Wei Lin and Mohammad T. Alhawary

2 Production of Modern Standard Arabic lexical stress cues by native

speakers of American English 38
Mashael Al-Aloula

3 Native English speakers’ perception and production of Arabic

consonants 56
Asmaa Shehata

4 The perception and production of Arabic consonants: a cross-

linguistic study 70
Sara Al Tubuly

5 Arabic L2 phonological acquisition: an ultrasound study of

emphatics and gutturals 93
Amanda Eads, Jodi Khater, and Jeff Mielke

v
 Contents

6 The L2 acquisition of Modern Standard Arabic final consonant

clusters by L1 Chinese speakers 113
Mona Maamoun

PART II
Arabic L2 vocabulary 137

7 Looking at words: an eye-tracking investigation of L2 Arabic

vocabulary learning 139
Ayman A. Mohamed

8 Keyword vs. context strategies among different levels of Arabic

language learners 157
Olla Najah Al-Shalchi

PART III
Arabic L2 morphosyntax 179

9 The acquisition of resumptive pronouns: how do second language

learners of Arabic do it? 181
Dola Algady

10 Arabic L2 learners’ use of word order and subject-verb agreement

for actor role assignment 201
Jamil Al-Thawahrih

PART IV
Arabic L2 reading and corpus-aided language learning 223

11 Corpus linguistics and critical reading and thinking: proposals for

teaching learning sequences based on journalistic corpora in Modern
Standard Arabic 225
Nadia Makouar

PART V
Arabic L2 writing: discourse analysis and measuring production 249

12 Writing in Arabic: discourse analysis and pedagogical reflections 251

Dris Soulaimani

vi
 Contents

13 Comparing the complexity, accuracy, and fluency of written Arabic

in the production of advanced learners and native speakers 265
Michael Raish

PART VI
Arabic L2 speaking and intercultural learning (in study abroad) 287

14 Code-switching in L2 Arabic collaborative dyadic interactions 289

Khaled Al Masaeed

15 Research-based interventions for language and intercultural learning 303

Emma Trentman

PART VII
Arabic heritage learners 329

16 Proficiency in standard Arabic and its predictors: the case of heritage

speakers in college-level elementary Arabic classrooms 331
Abdulkafi Albirini

17 Effect of age of acquisition on concept mediation in heritage Arabic

bilinguals 362
Iyad Ghanim

PART VIII
The Arabic L2 teacher: teacher training and self-positioning 385

18 Effect of using a collaborative video-based self-evaluation activity

on helping AFL student-teachers tie theory to practice 387
Raghda El Essawi

19 Arabic language teaching in the U.S.: two Arabic language users’

views on culture and self-positioning as teachers 402
Brahim Oulbeid

Index422

vii
 CONTRIBUTORS

Mashael Al-Aloula, George Mason University, USA

Abdulkafi Albirini, Utah State University, USA

Dola Algady, Al-Zahra College for Women, Muscat, Oman

Mohammad T. Alhawary, University of Michigan, USA

Khaled Al Masaeed, Carnegie Mellon University, USA

Olla Najah Al-Shalchi, University of Texas, Austin, USA

Jamil Al-Thawahrih, Defense Language Institute, Monterey, USA

Sara Al Tubuly, Al Maktoum College of Higher Education, UK

Amanda Eads, North Carolina State University, USA

Raghda El Essawi, American University in Cairo, Egypt

Iyad Ghanim, Montclair State University, USA

Jodi Khater, North Carolina State University, USA

Cheng-Wei Lin, University of Michigan, USA

Mona Maamoun, Alexandria University, Egypt

Nadia Makouar, INALCO, Paris, France

Jeff Mielke, North Carolina State University, USA

viii
 Contributors

Ayman A. Mohamed, Michigan State University, USA

Brahim Oulbeid, University of Massachusetts Amherst, USA

Michael Raish, College of William and Mary, USA

Asmaa Shehata, University of Ottawa, Canada

Dris Soulaimani, San Diego State University, USA

Emma Trentman, The University of New Mexico, USA

ix
 ACKNOWLEDGMENTS

Numerous individuals and entities contributed to this Handbook in many ways. First, a debt of
gratitude is owed to the anonymous participants of the studies reported on in this volume from
different parts of the world, without whom the studies and the Handbook would not have hap-
pened. Second, I would like to express my gratitude to the authors for their dedication to and
enthusiasm for the project. I also appreciate their patience with my multiple queries and
requests for revisions. Third, I am extremely grateful to the anonymous reviewers of all the
chapters. Their generous contributions of time and expertise contributed in no small measure
to the quality of this volume. Fourth, my thanks are extended to the various co-sponsors at the
University of Michigan who supported the conference “Investigating Arabic Second Lan-
guage Acquisition: Empirical Findings and Trends” held at the University of Michigan, Ann
Arbor, on September 23–25, 2016. The conference served as a significant discussion forum
and allowed most contributors to the volume to present their research studies and receive feed-
back from the audience. Co-sponsors of the conference included: Department of Near Eastern
Studies, Department of Linguistics, Center for Middle Eastern and North African Studies,
International Institute, Institute for the Humanities, Rackham Graduate School, and University
of Michigan Office of Research. My thanks also go to Andrea Hartill at Routledge, who
believed in and supported this project with foresight and enthusiasm. Last but not least, I am
forever grateful to my wife and children for putting up with the time taken away from them to
complete this volume and for their tremendous support and enthusiasm.

x
 INTRODUCTION

The present Handbook has been compiled in the hope of filling a critical gap in Arabic second
language acquisition (SLA) research. The objectives are to offer a range of Arabic SLA studies
which adopt state of the art research methods and techniques (qualitative and quantitative) in
various L2 subfields, provide empirical data and findings on many language components and
processes of Arabic as an L2, and generate more research interest in second language acquisi-
tion (SLA) studies – an area which remains underinvestigated. Because such research findings
contribute to our understanding of how Arabic as an L2 is learned, the findings are also crucial
to informing second language pedagogy and other related areas, such as testing and curriculum
development.
The vast majority of studies have so far been conducted on Arabic morphosyntax (e.g.,
Alhawary 2009 and Forthcoming; for a detailed review of such studies, see Alhawary 2009,
pp. 21–48). Little has been investigated with respect to other L2 Arabic language components
(such as phonology/pronunciation, vocabulary, and interlanguage pragmatics) and the differ-
ent learning processes and skills (reading, writing, speaking, and listening), and only recently
do we find studies on Arabic heritage learners, although the latter studies are part of a recently
emerging subfield (for a detailed survey of such studies, see Alhawary 2018). It is hoped from
this point on that Arabic SLA will continue to investigate as well as expand investigation cov-
erage of the different Arabic language components and the different aspects related to the
processes of Arabic second language learning so that adequate research coverage (both in
depth and breadth) can be achieved. By the same token, it will also be important to explore
other emerging approaches such as usage-based and emergentist or input frequency accounts
(e.g., Bybee and Hopper 2001; Ellis 2002, 2012; Agren and Van de Weijer 2013; Rebuschat
and Williams 2012; Römer et al. 2014), beside established rule-based approaches, prominent
among which is the formal generative framework.
Since the field of Arabic SLA is generally underinvestigated, it made little sense to invite
contributors to write chapters exploring second language acquisition issues and approaches
from a general perspective. Rather, the intention is to focus on works which are specifically
conducted on Arabic SLA. This has the additional advantage of enriching the burgeoning Ara-
bic SLA field and providing empirical evidence on various aspects of second language devel-
opment. Furthermore, because of the close affinity between second language learning and

1
 Introduction

second language teaching and because understanding how Arabic as an L2 is learned can
crucially inform teaching it, I asked authors to provide, in the conclusion section of each chap-
ter, specific implications of how their study findings may be applied in instructional contexts
so that both the Arabic L2 learner and Arabic foreign language teacher may benefit from
research results.
The chapters included in this volume address issues and aspects in Arabic L2 phonology
and phonetics, vocabulary learning, morphosyntax, reading, writing, speaking, and Arabic
heritage learners, as well as the Arabic L2 teacher. The studies represent current trends and
approaches in the field of second language acquisition as applied to Arabic on a range of areas
of Arabic second language acquisition and second language development. The book is divided
into eight parts comprising 19 chapters.
The first part contains six chapters and deals with L2 Arabic phonology and phonetics. The
first two chapters by Lin and Alhawary and Al-Aloula investigate an area in L2 Arabic pho-
nology (suprasegmental features) which had been unexamined before: lexical stress. The first
chapter, by Lin and Alhawary, investigates the perception and production of lexical stress by
L1 English and L1 Chinese learners of Arabic and explores input frequency and L1 transfer
effects. The second chapter, by Al-Aloula, examines L1 transfer and the production of lexical
stress by L1 English learners of Arabic by focusing on the phonetic cues of duration, pitch,
and intensity. The third chapter, by Shehata, explores the ability of L1 English learners of
Arabic to perceive and produce 10 Arabic consonant contrasts and the relationship between
learners’ perception and production. The fourth chapter, by Al Tubuly, investigates the role of
L1 in the perception and production of Arabic emphatic, uvular, and pharyngeal consonants
by Arabic L2 learners who are L1 speakers of English, Greek, Chinese, Turkish, and German.
The fifth chapter, by Eads et al., is an ultrasound study of Arabic emphatic and guttural con-
sonants and provides a detailed analysis and exploration of how L1 English learners of Arabic
articulate them, at what level they are able to differentiate their articulation, and in what order.
The sixth chapter, by Maamoun, investigates the validity of the universal sonority scale prin-
ciple in acquiring the pronunciation of Arabic final consonant clusters by L1 Chinese learners
of Arabic and the difficulty such learners encounter with Arabic word-final consonant
clusters.
The second part includes two chapters which focus on L2 Arabic vocabulary. Chapter 7, by
Mohamed, is an eye-tracking study of L2 Arabic vocabulary learning by English-speaking
learners of Arabic and explores whether or not the amount of attention to (novel) words over
repeated encounters can predict readers’ incidental acquisition of multiple aspects of vocabu-
lary knowledge. Chapter 8, by Al-Shalchi, compares experimentally between two vocabulary
learning strategies (keyword and context) by L1 English learners of Arabic at various levels of
proficiency to determine the impact of each on vocabulary learning and perceived workload
during instruction.
The third part focuses on the acquisition of Arabic L2 morphosyntax. Chapter 9, by Algady,
presents a (generative) minimalist account of the role of economy conditions on the syntactic
derivation of Arabic L2 resumptive pronouns within four types of relative clause construc-
tions: direct object, indirect object, oblique, and subject relative clauses. Chapter 10, by Al-
Thawahrih, examines the competition model’s prediction of L2 learners’ initial reliance on L1
cues of word order in processing and interpreting sentence structure by L1 English learners of
Arabic and whether such learners would instead use an L2 gender verbal agreement cue in
assigning the actor role.
The fourth part relates primarily to Arabic L2 reading and the use of corpus linguistics and
includes one chapter (Chapter 11), by Makouar. The chapter extends the application of interpretive

2
 Introduction

semantics theory and use of corpus analysis tools to Arabic L2 reading as a framework which aims
at developing the leaner’s metalinguistic awareness of word meanings and textual analysis in order
to achieve deeper understanding of texts and for L2 learners to become critical readers. The
approach is demonstrated in a case study with four French-speaking learners of Arabic at the inter-
mediate proficiency level.
The fifth part focuses on Arabic L2 writing from a discourse analysis perspective as well as
measuring writing development. In Chapter 12, Soulaimani draws on a discourse analysis
framework, which employs in particular the notions of voices and stance, in analyzing writ-
ings by advanced English-speaking learners of Arabic for use of discursive features and strate-
gies and compares such L2 writing output production with that of L1 Arabic speakers.
Similarly, in Chapter 13, Raish examines writings by advanced English-speaking learners by
comparing them with those of native Arabic speakers and applying the CAF framework with
its tripartite statistical measure of complexity, accuracy, and fluency.
The sixth part deals with promoting Arabic L2 speaking skill and language and intercultural
learning. The two chapters included also happen to address speaking and language and cultural
learning in the context of study abroad. In Chapter 14, Al Masaeed follows a nuanced under-
standing of Myers-Scotton’s markedness model for code-switching and explores code-
switching (Arabic to English) functions during conversation dyads between English-speaking
Arabic L2 learners and their Arabic native-speaking language partners outside of the class-
room. In Chapter 15, Trentman reports (qualitatively) on two case studies of three English-
speaking learners of Arabic. One case study includes one participant who was pursuing an
ethnographic project for study abroad; the other study includes two participants who took part
in a classroom telecollaboration project, one as a student and the other as a language partner.
The two studies explore the impact of inclusion of “guided reflection” (that draws from lin-
guistic and cultural evidence) on the participants’ language and cultural learning during tel-
ecollaboration and study abroad, respectively.
The seventh part relates to Arabic heritage learners. In Chapter 16, Albirini analyzes the
oral and written production of 29 Arabic heritage learners in terms of the three proficiency
measures of fluency, (grammatical) accuracy, and (syntactic) complexity. Based on the partici-
pants’ functional command in standard Arabic, Albirini discusses why it is theoretically and
pedagogically crucial not to approach heritage speakers on a par with foreign/second language
learners of Arabic. In Chapter 17, Ghanim investigates the implications of a lexico-semantic
model of bilingual lexical storage and retrieval and whether it is no longer possible for heritage
speakers (as a population of bilinguals who learned one language, Arabic, in childhood but lost
fluency on their way to adulthood) to activate the semantic information of (Arabic) lexical
items directly from the conceptual (mental) domain, and whether instead they do so (indi-
rectly) from semantic information associated with words in the other language (English). To
test this prediction and to dissociate between age and proficiency effects, Ghanim employs a
picture-naming and a translation task with 11 heritage Arabic-English bilinguals with varying
degrees of Arabic proficiency.
The book concludes with Part VIII, especially devoted to the Arabic second/foreign lan-
guage teacher, due to the special emphasis of the present volume and stronger connection it
attempts to make between second language acquisition research and second language peda-
gogy and bring more alignment between the two so that effective language learning and teach-
ing can be achieved. More research on teacher preparation, teacher education, teacher
cognition, and other internal and external variables is needed to examine arising issues and
needs of the teacher so that they can be understood and better met. Some of these aspects are
investigated in Chapters 18–19. Chapter 18, by El Essawi, is an evaluative study of

3
 Introduction

student-teachers’ level of self-reflection following a collaborative activity which employed the

Vialogues video discussion tool. Self-reflection in the study is defined as the ability to draw
upon abstract theoretical knowledge gained in teacher-education and teacher-training pro-
grams in order to evaluate the effectiveness of student-teachers’ own pedagogical practices.
The study focuses on five student-teacher participants who were native speakers of Arabic.
Chapter 19, by Oulbeid, is a case study of two non-native Arabic speaker college-level teach-
ers. It starts by exploring their beliefs about Arab culture and how such beliefs affect their
classroom practices, then investigates how the two teachers negotiate and construct their iden-
tities as teachers of Arabic as a foreign language and culture professionals and how they cat-
egorize themselves as non-native speaking teachers.
The Handbook will serve as a vital resource for Arabic SLA researchers and students, Ara-
bic applied linguists, Arabic teachers and practitioners of Arabic foreign language teaching,
Arabic language testers, and curriculum developers. It can be used as a course book in any
courses related to Arabic applied linguistics, Arabic second language acquisition, and foreign
language Arabic teaching methodology by graduate students and upper undergraduate stu-
dents. The Handbook will also be useful to scholars and practitioners in second language
acquisition and other neighboring disciplines, including theoretical linguistics, psycholinguis-
tics, and cognitive psychology.

Transcription/transliteration symbols
The transcription of all Arabic texts in the body of the Handbook follows a uniform system of
simplified and slightly modified International Phonetic Alphabet (IPA) symbols, with some
standard equivalents used widely in Arabic and Middle Eastern studies journals. For the pur-
pose of the chapters included in the volume, Table 0.1 represents the phonetic chart of Arabic
(MSA) consonants and their symbols used throughout. As for vowels, Table 0.2 lists MSA
vowels and some variants of dialectal varieties as cited in the chapters. For the citation of
Arabic titles and names of Arab authors in the references sections, a simplified transliteration
system based on standard usage in Arabic and Middle Eastern studies journals has been
adopted. The symbol ’ represents the hamza (glottal stop) and ‘ represents the ‘ayn (voiced
pharyngeal fricative consonant).

4
Table 0.1 Arabic (MSA) phonetic chart of consonants and their symbols

Arabic Labial Plain Emphatic Palato- Palatal Velar Uvular Pharyngeal Glottal
Consonants alveolar

Dental

Alveolar

Dental

Alveolar
Stop b t d ṭ ḍ k q ʔ
‫ب‬ ‫ت‬ ‫د‬ ‫ض ط‬ ‫ك‬ ‫ق‬ ‫ء‬
Nasal (Stop) m n
‫م‬ ‫ن‬
Fricative f θ ð s z ð ṣ š x γ ħ ʕ h
‫ف‬ ‫ث‬ ‫س ذ‬ ‫ظ ز‬ ‫ص‬ ‫ش‬ ‫خ‬ ‫ح غ‬ ‫هـ ع‬
Affricate ž
‫ج‬
Tap/Trill r
‫ر‬
Lateral l
‫ل‬
Glide w y w
‫و‬ ‫ي‬ ‫و‬
By convention (see also Ladefoged 2001), consonants to the left of a cell are voiceless and those to the
right are voiced; note also, in particular, /d/ and /ḍ/ are not complete (plain versus emphatic) contrasts,
since the production of /ḍ/ involves additionally both sides of the tongue touching/pressing against the
inner sides of the upper molars. In addition, depending on their own dialectal varieties, Arabic native
speakers may produce the uvular fricatives as velar fricatives.

Table 0.2 Arabic vowels (for broad transcription)

Arabic Symbol Transliteration Symbol Description

a short front/back low

‫ا‬ aa long front/back low

u short high back rounded

‫و‬ uu long high back rounded

i short high front unrounded (MSA)

e short mid front unrounded (colloquial)

‫ي‬ ii long high front unrounded

ْ‫ي‬ ay diphthong (MSA)

ee monophthong (colloquial)

‫ْو‬ aw diphthong (MSA)

oo monophthong (colloquial)
 Introduction

References
Agren, M. and Van de Weijer, J., 2013. Input frequency and the acquisition of subject-verb agreement in
number in spoken and written French. Journal of French Language Studies, 23 (3), 311–333.
Alhawary, Mohammad T., 2009. Arabic second language acquisition of morphosyntax. New Haven, CT:
Yale University Press.
Alhawary, Mohammad T., 2018. Empirical directions in the future of Arabic second language acquisition
and second language pedagogy. In: Kassem M. Wahba, et al., eds. Handbook for Arabic language
teaching professionals in the 21st century. New York: Routledge, 408–421.
Alhawary, Mohammad T., Forthcoming. Arabic second language learning and effects of input, transfer,
and typology. Washington, DC: Georgetown University Press.
Bybee, J. L. and Hopper, P. J., 2001. Frequency and the emergence of linguistic structure. Amsterdam:
John Benjamins.
Ellis, N. C., 2002. Frequency effects in language acquisition: A review with implications for theories of
implicit and explicit language acquisition. Studies in Second Language Acquisition, 24 (2), 143–188.
Ellis, N. C., 2012. Frequency-based accounts of second language acquisition. In: S. Gass and A. Mackey,
eds. The Routledge handbook of second language acquisition. New York: Routledge, 193–210.
Ladefoged, Peter. 2001. A course in phonetics. Boston, MA: Heinle and Heinle.
Rebuschat, P. and Williams, J. N. 2012. Statistical learning and language acquisition. Berlin: De Gruyter
Mouton.
Römer, U., et al., 2014. Second language learner knowledge of verb-argument constructions: Effects of
language transfer and typology. The Modern Language Journal, 98 (4), 952–975.

6
 Cheng-Wei Lin and Mohammad T. Alhawary

1.4.3.1 Stress identification task and procedure

The purpose of the stress identification task was to generally investigate learners’ perception
of stress and whether their perception was influenced by the different frequency cues within
the stimuli. In this task, the stimuli were presented to participants both aurally and visually. For
each token, participants were presented with a recording of the stimulus produced by a male
native speaker of Arabic (from Saudi Arabia) and saw the orthography of the word at the same
time on the computer screen. After listening to the word, participants were asked to identify
the stressed syllable in a word by pressing the corresponding number on the first row of the
control panel, where the number “1” stood for the first syllable, “2” stood for the second syl-
lable, and “3” for the third syllable. The participants were asked to complete each token as
soon as they could.
Since participants might not know the syllabification of the stimuli solely based on Arabic
orthography, the orthography of the word was syllabified and numbered, but vowel informa-
tion (short and long) was eliminated from the visual stimuli, as in the word ‫ نتيجة‬na.ˈtiː.ža
“result” appearing without vowels, as shown in Figure 1.5, so that the participants could only
be shown that the stimulus had three syllable: with [n] being the onset of the first syllable, [t]
the second syllable, and [ž] the third. This was deemed necessary in order to avoid making the
orthography reveal the pronunciation of a word and making participants rely on the visual
aspect of the stimuli alone when identifying the stressed syllable, since Arabic spelling (with
vowels included) is transparent, unlike English. Another reason for concealing vowel informa-
tion was to avoid scenarios where learners might impressionistically but falsely prefer a cer-
tain type of vowel quality or vowel length in the script of stimuli without attending to the
acoustic cues provided by the (aural) stimuli.

1.4.3.2 Results of stress identification task

Both the fluency and the accuracy of participants’ responses to the stimuli were analyzed. Flu-
ency in this task and throughout is operationalized as the reaction time or amount of time the
participants took to respond to the stimuli. Accuracy refers to whether the syllable selected by
the participants matches the attested position for the stressed syllable in the language, and
responses are accordingly marked as correct or incorrect. Both measures provide different
dimensions of assessing participants’ performance in stress perception: their performance was
deemed worse if it took them a longer time to figure out the stress position, even if they did so
correctly. Similarly, no matter how fast their response time was, their performance was consid-
ered worse if they did not identify the position of the stressed syllable correctly. Equally
important here is that only participants’ response to stimuli that were nonsense words were
analyzed, to avoid bias introduced from familiarity effects.
To examine the effect of L1 on reaction time, a two-way ANOVA was conducted. The
analysis revealed a main effect for L1 (F(2, 73) = 4.98, p < 0.001, partial η2 = 0.11) but no
proficiency or interaction effect was found. Follow-up post hoc comparisons using Tukey tests

Figure 1.5 Sample of stimuli presentation in the stress identification task

18
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globules, these latter being the fatty portion of the yolk; these are present
everywhere, though in the centre there is a space where they are very
scanty, and they also do not extend quite to the circumference. But the
most remarkable change that has taken place is the appearance in the
middle of the field of an area different from the rest in several particulars;
it occupies about one-third of the width and one-third of the length; it has
a whiter and more opaque appearance, and the fat globules in it are
fewer in number and more indistinct. This area is afterwards seen to be
occupied by the developing embryo, the outlines of which become
gradually more distinct. Fig. 83 gives an idea of the appearance of the
egg about the middle period of the development. In warm weather the
larva emerges from this egg ten or eleven days after it has been
deposited.
 Fig. 83.—A, Egg of Limacodes testudo about the middle of the development
of the embryo; B, micropyles and surrounding sculpture of chorion.

The period occupied by the development of the embryo is very different in

the various kinds of Insects; the blowfly embryo is fully developed in less
than twenty-four hours, while in some of the Orthoptera the embryonic
stage may be prolonged through several months. According to
Woodworth the blastoderm in Vanessa antiopa is complete in twenty-four
hours after the deposition of the egg, and the involution of the ventral
plate is accomplished within three days of deposition.

Metamorphosis.

The ontogeny, or life history of the individual, of Insects is peculiar,

inasmuch as a very large part of the development takes place only late in
life and after growth has been completed. Insects leave the egg in a
certain form, and in that condition they continue—with, however, a greater
or less amount of change according to kind—till growth is completed,
when, in many cases, a very great change of form takes place. Post-
embryonic development, or change of form of this kind, is called
metamorphosis. It is not a phenomenon peculiar to Insects, but exists to a
greater or less extent in other groups of the Metazoa; while simpler post-
embryonic development occurs in nearly all, as in scarcely any complex
animals are all the organs completely formed at the time the individual
becomes possessed of a separate existence. In many animals other than
Insects the post-embryonic development assumes most remarkable and
complex forms, though there are perhaps none in which the phenomenon
is very similar to the metamorphosis of Insects. The essential features of
metamorphosis, as exhibited in the great class we are writing of, appear
to be the separation in time of growth and development, and the limitation
of the reproductive processes to a short period at the end of the individual
life. The peculiar phenomena of the post-embryonic development of the
white ants show that there exists some remarkable correlation between
the condition of the reproductive organs and the development of the other
parts of the organisation. If we take it that the post-embryonic
physiological processes of any individual Insect are of three kinds,—
growth, development, and reproduction,—then we may say that in the
higher Insects these three processes are almost completely separated,
and go on consecutively, the order being,—first, growth; second,
development; third, reproduction. While, if we complete the view by
including the processes comprised in the formation of the egg and the
development therein, the series will be—(1) oogenesis, or egg-growth; (2)
development (embryonic); (3) growth (post-embryonic); (4) development
(post-embryonic); (5) reproduction.

The metamorphosis of Insects is one of the most interesting parts of

entomology. It is, however, as yet very little known from a scientific point
of view, although the simpler of its external characters have for many
ages past attracted the attention and elicited the admiration of lovers of
nature. It may seem incorrect to say that little is yet known scientifically of
a phenomenon concerning which references almost innumerable are to
be found in literature: nevertheless the observations that have been made
as to metamorphosis, and the analysis that has been commenced of the
facts are at present little more than sufficient to show us how vast and
complex is the subject, and how great are the difficulties it presents.

There are three great fields of inquiry in regard to metamorphosis, viz. (1)
the external form at the different stages; (2) the internal organs and their
changes; (3) the physiological processes. Of these only the first has yet
received any extensive attention, though it is the third that precedes or
underlies the other two, and is the most important. We will say a few
words about each of these departments of the inquiry. Taking first the
external form—the instar. But before turning to this we must point out that
in limiting the inquiry to the post-embryonic development, we are making
one of those limitations that give rise to much misconception, though they
are necessary for the acquisition of knowledge as to any complex set of
phenomena. If we assume five well-marked stages as constituting the life
of an Insect with extreme metamorphosis, viz. (1) the formation and
growth of the egg; (2) the changes in the egg culminating in its hatching
after fertilisation; (3) the period of growth; (4) the pupal changes; (5) the
life of the perfect Insect; and if we limit our inquiry about development to
the latter three, we are then shutting out of view a great preliminary
question, viz. whether some Insects leave the egg in a different stage of
development to others, and we are consequently exposing ourselves to
the risk of forgetting that some of the distinctions we observe in the
subsequent metamorphosis may be consequential on differences in the
embryonic development.
 Instar and Stadium.

Figs. 84 and 85 represent corresponding stages in the life of two different

Insects, Fig. 84 showing a locust (Acridium), and Fig. 85 a white butterfly.
In each A represents the newly-hatched individual; B, the insect just
before its perfect state; C, the perfect or imago stage. On comparing the
two sets of figures we see that the C stages correspond pretty well as
regards the most important features (the position of the wings being
unimportant), that the A stages are moderately different, while the B
states are not to be recognised as equivalent conditions.

Fig. 84.—Locust (Acridium peregrinum): A, newly hatched; B, just

antecedent to last ecdysis; C, perfect Insect.

Fig. 85.—Butterfly (Pieris): A, the newly hatched young, or larva magnified;

B, pupa (natural size) just antecedent to last ecdysis; C, perfect Insect.

Every Insect after leaving the egg undergoes during the process of
growth castings of the skin, each of which is called a moult or ecdysis.
Taking for our present purpose five as the number of ecdyses undergone
by both the locust and butterfly, we may express the differences in the
successions of change we portray in Figs. 84 and 85 by saying that
previous to the first ecdysis the two Insects are moderately dissimilar, that
the locust undergoes a moderate change before reaching the fifth
ecdysis, and undergoes another moderate change at this moult, thus
reaching its perfect condition by a slight, rather gradual series of
alterations of form. On the other hand, the butterfly undergoes but little
modification, remaining much in the condition shown by A, Fig. 85, till the
fourth, or penultimate, ecdysis, but then suffers a complete change of
form and condition, which apparently is only inferior to another
astonishing change that takes place at the fifth or final moult. The chief,
though by no means the only, difference between the two series consists
in the fact that the butterfly has interposed between the penultimate and
the final ecdyses a completely quiescent helpless condition, in which it is
deprived of external organs of sense, locomotion, and nutrition; while in
the locust there is no loss of these organs, and such quiescent period as
exists is confined to a short period just at the fifth ecdysis. The changes
exhibited by the butterfly are called "complete metamorphosis," while this
phenomenon in the locust is said to be "incomplete." The Insect with
complete metamorphosis is in its early stage called a larva, and in the
quiescent state a pupa. The adult state in both butterfly and locust is
known as imago or perfect Insect.

The most conspicuous of the differences between Insects with complete

and those with incomplete metamorphosis is, as we have remarked, the
existence in the former of a pupa. The pupal state is by no means similar
in all the Insects that possess it. The most anomalous conditions in
regard to it occur in the Order Neuroptera. In some members of that
Order—the Caddis-flies for instance—the pupa is at first quiescent, but
becomes active before the last ecdysis; while in another division—the
May-flies—the last ecdysis is not preceded by a formed pupa, nor is there
even a distinct pupal period, but the penultimate ecdysis is accompanied
by a change of form to the winged condition, the final ecdysis being
merely a casting of the skin after the winged state has been assumed. In
the Odonata or Dragon-flies there is no pupal stage, but the change of
form occurring at the last ecdysis is very great. In those Insects where the
interval between the last two moults is not accompanied by the creature's
passing into a definite, quiescent pupa, the individual is frequently called
then a nymph; but the term nymph has merely a distinctive meaning, and
is not capable of accurate definition, owing to the variety of different
conditions covered by the word. Eaton, in describing this term as it is
used for Ephemeridae, says, "Nymphs are young which lead an active
life, quitting the egg at a tolerably advanced stage of morphological
development, and having the mouth-parts formed after the same main
type of construction as those of the adult insect."[80]

The intervals between the ecdyses are called stadia, the first stadium
being the period between hatching and the first ecdysis. Unfortunately no
term is in general use to express the form of the Insect at the various
stadia; entomologists say, "the form assumed at the first moult," and so
on. To avoid this circumlocution it may be well to adopt a term suggested
by Fischer,[81] and call the Insect as it appears at hatching the first instar,
what it is as it emerges from the first ecdysis the second instar, and so on;
in that case the pupa of a Lepidopteron that assumed that condition at the
fifth ecdysis would be the sixth instar, and the butterfly itself would be the
seventh instar.

Various terms are used to express the differences that exist in the
metamorphoses of Insects, and as these terms refer chiefly to the
changes in the outer form, we will here mention them. As already stated,
the locust is, in our own language, said to have an incomplete
metamorphosis, the butterfly a complete one. The term Holometabola has
been proposed for Insects with complete metamorphosis, while the
appellations Ametabola, Hemimetabola, Heterometabola, and
Paurometabola have been invented for the various forms of incomplete,
or rather less complex, metamorphosis. Some writers use the term
Ametabola for Insects that are supposed to exhibit no change of external
form after quitting the egg, the contrasted series of all other Insects being
then called Metabola. Westwood and others use the word Homomorpha
for Insects in which the condition on hatching more or less resembles that
attained at the close of the development, and Heteromorpha for those in
which the form on emergence from the egg differs much from what it
ultimately becomes.

Hypermetamorphosis.

There are certain unusual changes to which the term

hypermetamorphosis has been applied; these we can here only briefly
allude to.
Insects that have complete metamorphoses, and are not supplied with
food by their parents or guardians, are provided during their larval life with
special modifications of extremely various kinds to fit them for the period
of life during which they are obtaining food and growing. Thus caterpillars
possess numerous adaptations to fit them for the period during which
they live on leaves, while maggots have modifications enabling them to
live amongst decomposing flesh. Some larvae are greatly modified in this
adaptive way, and when the adaptations change greatly during the life of
the larva, hypermetamorphosis is said to exist. As an instance we may
mention some beetle larvae that are born with legs by whose aid they can
cling to a bee, and so get carried to its nest, where they will in future live
on the stores of food the bee provides for its own young. In order that
they may be accommodated to their totally different second
circumstances, they change their first form, losing their legs, and
becoming almost bladder-like creatures, fitted for floating on the honey
without being injured by it. Such an occurrence has been described by
Fabre[82] in the case of Sitaris humeralis, and his figures have been
reproduced in Sir John Lubbock's book on the metamorphoses of Insects,
[83] as well as in other works, yet they are of so much interest that we give
them again, especially as the subject is still only in its infancy; we at
present see no sufficient reason for the later of these larval states. Little
is, we believe, known as to the internal anatomy of the various instars in
these curious cases.

Fig. 86.—Preparatory stages of Sitaris humeralis: 9, 10, 11, 12, first,

second, third, and fourth larval instars; 13, pupa. (After Lubbock and
Fabre.)

There are certain minute Hymenoptera that deposit their eggs inside the
eggs of other Insects, where the beings hatched from the parasitic eggs
subsequently undergo their development and growth, finding their
sustenance in the yolk or embryo contained in the host-egg. It is evident
that such a life is very anomalous as regards both food and the conditions
for respiration, and we consequently find that these tiny egg-parasites go
through a series of changes of form of a most remarkable character.[84] It
would appear that in these cases the embryonic and post-embryonic
developments are not separated in the same way as they are in other
Insects. We are not aware that any term has yet been proposed for this
very curious kind of Insect development, which, as pointed out by Brauer,
[85] is doubtless of a different nature from the hypermetamorphosis of
Sitaris.

Changes in Internal Organs.

In relation to the post-embryonic development of the internal organs of

the body there is but little exact generalisation to be made, the anatomical
condition of these organs at the time of emergence from the egg having
been ascertained in but few Insects. We know that in Holometabolous
Insects the internal anatomy differs profoundly in the larval and imaginal
instars. As to Insects with more imperfect metamorphosis very little
information exists, but it appears probable that in many no extensive
distinctions exist between the newly-hatched and the adult forms, except
in the condition of the reproductive organs. Differences of minor
importance doubtless exist, but there is almost no information as to their
extent, or as to the periods at which the changes occur; so that we do not
know to what extent they may be concentrated at the final ecdysis. In
Insects with perfect metamorphosis the structures of the internal organs
are, as we have said, in many cases totally different in the larval and
imaginal periods of the life; but these changes are far from being uniform
in all Holometabola. The nervous system in some cases undergoes a
great concentration of the ganglia, in others does not, and important
distinctions exist in this respect even within the limits of a single Order,
such as the Coleoptera. Some Insects take the same kind of food
throughout their lives, but many others change totally in this respect, and
their organs for the prehension and digestion of food undergo a
corresponding change. Butterflies suck food in the form of liquid juices
from flowers by means of a delicate and long proboscis, while the young
butterfly—the caterpillar—disdains sweets, and consumes, by the
assistance of powerful mandibles, a great bulk of leaves. Other
Holometabola undergo no such total change of habits; the tiger-beetle, for
instance, is as ferocious a consumer of the juices of Insects in its young
stage as it is in the adult condition. Hence Brauer[86] divides Insects, as
regards this point, into three categories. The forms in which both the
young and adult take food by suction he calls Menorhyncha; those in
which both the imago and immature forms feed by mandibles he calls
Menognatha; while his Metagnatha consists of those insects that take
food by jaws when young, but by suction with tubular mouths when
mature. Besides these main divisions there are some exceptional cases
to which we need not here allude, our present object being to indicate that
in the Metagnatha the digestive organs are of a very different nature in
the young and in the adult states of existence.

The internal organs for the continuance of the species are known to be
present in a rudimentary stage in the embryo, and it is a rule that they do
not attain their full development until growth has been completed; to this
rule there may possibly be an exception in the case of the Aptera. But
little information of a comparative character exists as to the dorsal vessel
and the changes it undergoes during metamorphosis. There is
considerable difficulty in connexion with the examination of this structure,
but it appears probable that it is one of the organs that changes the least
during the process of metamorphosis.

The exact nature of the internal changes that occur during

metamorphosis is almost a modern subject. It is of course a matter of
great difficulty to observe and record changes that go on in the interior of
such small creatures as Insects, and when the phenomena occur with
great rapidity, as is frequently the case in Insect metamorphosis, the
difficulty is much increased. Nevertheless the subject is of such great
interest that it has been investigated with a skill and perseverance that
call for the highest admiration. The greater part of the information
obtained refers to a single Insect, the blowfly; and amongst those who
have made important contributions to it we may mention Weismann,[87]
Viallanes,[88] Ganin,[89] and Van Rees,[90] and it is at present under
investigation by Lowne. A good deal, too, is becoming known about the
processes in the case of the silkworm.

Integument and Ecdysis.

The integument consists of a cellular layer, usually called the hypodermis,
situated on a basement membrane. The hypodermis, or layer of
chitinogenous cells, excretes a matter which remains attached to the
body, forming the hard outer layer of the skin. This layer consists of chitin
and has no vitality, but its presence no doubt exerts a very important
influence on the physiological processes of the Insect. The chitinous
investment varies much in thickness and in other properties; in some
Insects it is hard, even glassy, so as to be difficult to pierce with a pin, in
others it is pliable, and in some very delicate. Chitin is a substance very
difficult to investigate; according to the recent researches of Krawkow[91]
it may prove to be of somewhat variable chemical composition.

After a time the hypodermis excretes a fresh supply of chitin, and,

possibly by the commencement of this process, the older chitinous
investment becomes separated and is shed. The details have, however,
not been ascertained, though their importance has been suggested by
Hatchett Jackson.[92] The newly exposed layer of integument is pallid, but
afterwards becomes coloured in a manner varying according to the
species, the process being possibly due to some secondary exudation
permeating the freshly exposed chitin, or modifying some part of its
exterior.

Lowne informs us that in the imago of the blowfly the great majority of the
hypodermic cells themselves enter into the composition of the chitinous
integument; and it is perhaps not a matter for surprise that the cells
should die on the completion of their functional activity, and should form a
part of the chitinous investment. Some writers say that the chitinous layer
may be shown to be covered by a delicate extima or outer coat.

The number of ecdyses varies greatly in Insects, but has been definitely
ascertained in only a few forms outside the Order Lepidoptera. In
Campodea Grassi says there is a single fragmentary moult, and in many
Hymenoptera the skin that is cast is extremely delicate, and the process
perhaps only occurs twice or three times previous to the pupal stage. In
most Insects, however, ecdysis is a much more important affair, and the
whole of the chitinous integument is cast off entire, even the linings of the
tracheae, and of the alimentary canal and its adjuncts being parted with.
Sir John Lubbock observed twenty-three moults in a May-fly of the genus
Cloëon,[93] this being the maximum yet recorded, though Sommer
states[94] that in Macrotoma plumbea moulting goes on as long as life
lasts, even after the Insect has attained its full size.

Some Insects get quit of a considerable quantity of matter by their

ecdyses, while in others the amount is comparatively slight. It has been
thought that the moulting is effected in order to permit of increase of size
of the Insect, but there are facts which point to the conclusion that this is
only a factor of secondary importance in the matter. One of these is that
many Insects make their first ecdysis almost immediately after they leave
the egg; this is the case with the young larva of the blowfly, which,
according to Lowne, moults within two hours of its emergence from the
egg. We have already referred to the important suggestion made by
Eisig[95] that, since chitin is a nitrogenous substance, the ecdyses may be
a means of getting rid of waste nitrogenous matter; to which we have
added that as chitin also consists largely of carbon, its excretion may be
of importance in separating carbonaceous products from the blood.

Metamorphosis of Blowfly.

The phenomena of metamorphosis are displayed to their greatest extent

in the transformations and physiological processes of the Muscid Diptera,
of which the common blowfly is an example. We will briefly consider the
information that has been obtained on this subject.

The development of the embryo in the egg of the blowfly is unusually

rapid, occupying only a period of twenty to twenty-four hours. After its first
moult the blowfly larva grows rapidly during a period of about ten to
fourteen days, during which it undergoes moults, the number of which
appears not to be definitely ascertained. After becoming full-fed the larva
loses its active state, and passes for a period into a condition of
comparative quiescence, being spoken of in this state as a resting larva.
This quiet period occurs in most full-grown larvae, and is remarkable for
the great variation that may occur in its duration, it being in many Insects
subject to prolongation for months, in some cases possibly even for
years, though in favourable circumstances it may be very short. Lowne
informs us that in the blowfly this period of the life is occupied by very
great changes in the internal organs, which are undergoing very
extensive processes of destruction and rebuilding. After some days the
outer skin of the resting larva shrivels, and is detached from the internal
living substances, round which it hardens and forms the sort of cocoon or
capsule that is so well known. This using of the cast larval skin as a
cocoon is, however, limited to certain of the two-winged flies, and perhaps
a few other Insects, and so must be considered an exceptional condition.
The capsule conceals from view a most remarkable state, known to the
old naturalist Réaumur as the "spheroidal condition," but called by more
recent writers the pronymph. The pronymphal state may be looked on as
being to a great extent a return of the animal to the condition of an egg,
the creature becoming an accumulation of soft creamy matter enclosed in
a delicate skin. This spheroidal condition, however, really begins in the
resting larva, and Van Rees and others think that the delicate membrane
enclosing the substance of the pronymph is really the hypodermis of the
integument of the larva. Although this seems probable, from the
resemblance this condition would in that case present to the phenomena
usual in ecdysis, it is not generally admitted, and there is much difficulty in
settling the point. Lowne is of a contrary opinion, looking on the limiting
membrane as a subsequent formation; he calls it the paraderm. The
process of forming the various organs goes on in the pronymph, till the
"nymph" has completed its development, the creature having then again
taken on a definite form which apparently corresponds to the pupa of
Hymenoptera. Great doubt, however, exists as to this equivalence, and
indeed as to any exact correspondence between the metamorphic stadia
of different Insects, a view which long since was expressed by Sir John
Lubbock[96] and Packard. The term nymph is used in this case not
because there is any resemblance to the condition similarly named in
Insects with less complete metamorphosis, but because the term pupa is
applied to the outer case together with the contained nymph. The
transformation of the nymph into the perfect blowfly occupies a period
very variable according to the temperature.

Histolysis.—The processes by which the internal organs of the maggot

are converted into those of the fly are of two kinds,—histolysis or breaking
down, histogenesis or building up, of tissue. The intermediary agents in
histolysis are phagocytes, cells similar to the leucocytes or white
corpuscles of the blood: the intermediary agents in histogenesis are
portions of tissue existing in the larval state incorporated with the different
organs, or preserving a connexion therewith even when they are to a
great extent separated therefrom. In this latter case they are called
imaginal discs, though Professor Miall prefers to term them imaginal
folds.[97] The two processes of histolysis and histogenesis, though to
some extent mutually dependent (for the material to be built up has to be
largely obtained by previous destruction), do not go on pari passu, though
they are to a great extent contemporaneous. In the resting larva histolysis
is predominant, while in the nymph histogenesis is more extensive.
Microscopic observation shows that the phenomena connected with the
histolysis of the muscular tissue are scarcely distinguishable from those
of an inflammatory process, and Viallanes[98] dilates on this fact in an
instructive manner. The phagocytes attach themselves to, or enter, the
tissues which are to be disintegrated, and becoming distended, assume a
granular appearance. By this pseudo-inflammatory process the larval
structures are broken down into a creamy substance; the buds, or germs,
from which the new organs are to be developed being exempt from the
destruction. These buds, of which about sixty or upwards have already
been detected, undergo growth as they are liberated, and so the new
creature is formed, the process of growth in certain parts going on while
destruction is being accomplished in others. Considerable discrepancy
prevails as to the extent to which the disintegration of some of the tissues
is carried.

Fig. 87.—Imaginal discs of Muscidae in process of development: A, Brain

and ventral ganglion of a larva 7 mm. long of M. vomitoria; v, ventral
ganglion; c, cephalic ganglion; h, head rudiment; vc, portion of ventral
chain; pd, prothoracic rudiment; vc3, third nerve; md, mesothoracic
rudiment: B, mesothoracic rudiment, more advanced, in a pupa just
formed of Sarcophaga carnaria, showing the base of the sternum and
folds of the forming leg, the central part (f) representing the foot: C, the
rudimentary leg of the same more advanced; f, femur; t, tibia; f1, f5,
tarsal joints: D, two discs from a larva 20 mm. long of Sarcophaga,
attached to tracheae; msw, mesonotal and wing-rudiment; mt,
 metathoracic rudiment: E, r, mesothoracic rudiment of a 7 mm. long
larva attached to a tracheal twig. (After Weismann and Graber.)

According to Kowalevsky[99] it would appear that after the phagocytes

have become loaded with granules they serve as nutriment for the
growing tissues, and he thinks they become blood-cells in the imago. The
process of histolysis has been chiefly studied in the blowfly, and not much
is known of it in other Insects, yet it occurs to a considerable extent,
according to Bugnion[100] and others, in the metamorphosis of
Lepidoptera. Indeed it would almost seem that the processes of histolysis
and histogenesis may be looked on as exaggerated forms of the
phenomena of the ordinary life of tissues, due to greater rapidity and
discontinuity of tissue nutrition.

Imaginal Discs.—The imaginal discs are portions of the larval hypoderm,

detached from continuity with the main body of the integument, but
connected therewith by strings or pedicels which may be looked on as
portions of the basement membrane. Whether these discs, or histoblasts
as they are called by Künckel d'Herculais,[101] are distinguished by any
important character from other buds or portions of regenerative tissue
that, according to Kowalevsky,[102] Korschelt and Heider,[103] and others,
exist in other parts of the body, does not appear to be at present
ascertained.

We give some figures, taken from Weismann and Graber, of the imaginal
rudiments existing in the larvae of Muscidae. Although by no means
good, they are the best for our purpose we can offer to the reader. Other
figures will be found in Lowne's work on the blowfly now in course of
publication. Weismann's paper[104] is now thirty years old, and, when it
was written, he was not aware of the intimate connexion the rudiments
have with the integument; this has, however, now been demonstrated by
several observers. Pratt states[105] that the formation of the imaginal
discs in Melophagus ovinus takes place in the later stages of the
embryonic development, and after the manner formerly suggested by
Balfour, viz. invagination of the ectoderm.
 Fig. 88.—Median longitudinal section through larva of blowfly during the
process of histolysis. (After Graber.) Explanation in text.

Both the regenerative buds and the rudimentary sexual glands are known
to be derived directly from the embryo; neither of them undergoes any
histolysis, so that we have in them embryonic structures which exist in a
quiescent condition during the period in which the larva is growing with
great rapidity, and which when the larva has attained its full growth and is
disintegrating, then appropriate the products of the disintegration so as to
produce the perfect fly.

Our Fig. 88, taken from Graber, represents a longitudinal median section
of a full-grown larva of Musca, in which the processes of metamorphosis
are taking place. The position of some of the more important imaginal
rudiments is shown by it: b1, b2, b3, rudiments of the three pairs of legs of
the imago; an, of antennae; between an and w, rudiment of eye; w, of
wings; h, of halteres; f, fat-body; d, middle of alimentary canal; n, ventral
chain; st, stigma; 6, 7, sixth and seventh body segments.

Physiology of Metamorphosis.

Many years ago, Harvey perceived the probable existence of a

physiological continuity between the earlier and later stages of the
Insect's life. Modern investigation has shown that in the blowfly a
remarkable analogy exists between the conditions of the pupa and the
egg. The outer shell of the pupa corresponds to the chorion or egg-shell,
and the delicate outer membrane of the pronymph to the oolemn or lining
membrane of the egg; the creamy matter corresponds with the yolk, and
the regenerative buds are analogous to the formative portions of the
developing egg. The process of histolysis as carried out by the
phagocytes of the later life appears also to find a parallel in the
vitellophags of the embryonic life.[106] It appears probable that the
physiological processes of the post-embryonic metamorphosis may be
essentially a repetition—or an interrupted continuation—of those of the
embryonic period.

The inquiry as to what are the determining causes of the metamorphic

changes of the blowfly and other Insects has as yet but little advanced.
Why does the larva grow up to a certain period with great rapidity, then
cease its appropriating power and break up the parts that have been so
rapidly and recently formed? And why do the imaginal buds remain
quiescent till the other tissues are being disintegrated, and then, instead
of sharing the general condition of disintegration, commence a career of
development? To these questions no satisfactory answer has yet been
given, though the remarkable studies, already referred to, of Bataillon on
the later larval life of the silkworm suggest the direction in which
knowledge may be found, for they show that the physiological conditions
of the later larval life are different from those of the earlier life, possibly as
the direct result of the mere aggregation of matter, and the consequent
different relations of the parts of the organism to atmospheric and
aqueous conditions.

If we wish to understand metamorphosis, we must supplement the old

opinion that ecdysis is merely an occurrence to facilitate expansion, by
the more modern conception that it is also an important physiological
process. That shedding the skin is done solely to permit of enlargement of
size is a view rendered untenable by many considerations. The
integument can increase and stretch to an enormous extent without the
aid of moulting; witness the queen-termite, and the honey-bearers of the
Myrmecocystus ants. Many moults are made when increase of size does
not demand them, and the shedding of the skin at the time of pupation is
accompanied by a decrease in size. And if moulting be merely connected
with increase of size, it is impossible to see why Cloëon should require
two dozen moults, while Campodea can do with one, or why a
collembolon should go on moulting during the period of life subsequent to
the cessation of growth.
The attention of entomologists has been chiefly directed to the ecdyses
connected with the disclosure of the pupal and imaginal instars. Various
important transformations may, however, occur previous to this, and when
they do so it is always in connexion with ecdyses. Caterpillars frequently
assume a different appearance and change their habits or character at a
particular ecdysis; and in Orthoptera each ecdysis is accompanied by a
change of form of the thoracic segments; this change is very considerable
at one of the intermediate ecdyses.

The assumption of the pupa state is the concomitant of an ecdysis, and

so also is the appearance of the imago; but the commencement of each
of these two stages precedes the ecdysis, which is merely the outward
mark of the physiological processes. The ecdysis by which the pupa is
revealed occurs after the completion of growth and when great changes
in the internal organs have occurred and are still taking place; the ecdysis
by which the imago appears comes after development has been quite or
nearly completed.

Although the existence of a pupa is to the eye the most striking of the
differences between Insects with perfect and those with imperfect
metamorphosis, yet there is reason for supposing that the pupa and the
pupal period are really of less importance than they at first sight appear to
be. In Fig. 85 we showed how great is the difference in appearance
between the pupa and the imago. The condition that precedes the
appearance of the pupa is, however, really the period of the most
important change. In Fig. 89 we represent the larva and pupa of a bee; it
will be seen that the difference between the two forms is very great, while
the further change that will be required to complete the perfect Insect is
but slight. When the last skin of the larva of a bee or of a beetle is thrown
off, it is, in fact, the imago that is revealed; the form thus displayed,
though colourless and soft, is that of the perfect Insect; what remains to
be done is a little shrinking of some parts and expansion of others, the
development of the colour, the hardening of certain parts. The colour
appears quite gradually and in a regular course, the eyes being usually
the first parts to darken. After the coloration is more or less perfected—
according to the species—a delicate pellicle is shed or rubbed off, and the
bee or beetle assumes its final form, though usually it does not become
active till after a farther period of repose.
 Fig. 89.—Larva and pupa of a bee, Xylocopa violacea: A, larva; B, pupa,
ventral aspect; C, pupa, dorsal aspect. (After Lucas.)

CHAPTER VI

CLASSIFICATION—THE NINE ORDERS OF INSECTS—THEIR CHARACTERS—

PACKARD'S ARRANGEMENT—BRAUER'S CLASSIFICATION—CLASSIFICATIONS
BASED ON METAMORPHOSIS—SUPER-ORDERS—THE SUBDIVISIONS OF
ORDERS.

Classification.

We have already alluded to the fact that Insects are the most numerous in
species and individuals of all land animals: it is estimated that about
250,000 species have been already described and have had scientific
names given to them, and it is considered that this is probably only about
one-tenth of those that really exist. The classification in a comprehensible
manner of such an enormous number of forms is, it will be readily
understood, a matter of great difficulty. Several methods or schemes have
since the time of Linnaeus been devised for the purpose, but we shall not
trouble the reader to consider them, because most of them have fallen
into disuse and have only a historical interest. Even at present there
exists, however, considerable diversity of opinion on the question of
classification, due in part to the fact that some naturalists take the
structure of the perfect or adult Insect as the basis of their arrangement,
while others prefer to treat the steps or processes by which the structure
is attained, as being of primary importance. To consider the relative
values of these two methods would be beyond our scope, but as in
practice a knowledge of the structures themselves must precede an
inquiry as to the phases of development by which the structures are
reached; and as this latter kind of knowledge has been obtained in the
case of a comparatively small portion of the known forms,—the
embryology and metamorphosis having been investigated in but few
Insects,—it is clear that a classification on the basis of structure is the
only one that can be at present of practical value. We shall therefore for
the purposes of this work make use of an old and simple system, taking
as of primary importance the nature of the organs of flight, and of the
appendages for the introduction of food to the body by the perfect Insect.
We do not attempt to disguise the fact that this method is open to most
serious objections, but we believe that it is nevertheless at present the
most simple and useful one, and is likely to remain such, at any rate as
long as knowledge of development is in process of attainment.

Orders.

The great groups of Insects are called Orders, and of these we recognise
nine, viz. (1) Aptera, (2) Orthoptera, (3) Neuroptera, (4) Hymenoptera, (5)
Coleoptera, (6) Lepidoptera, (7) Diptera, (8) Thysanoptera, (9) Hemiptera.
These names are framed to represent the nature of the wings; and there
is some advantage in having the Orders named in a uniform and
descriptive manner. The system we adopt differs but little from that
proposed by Linnaeus.[107] The great Swedish naturalist did not,
however, recognise the Orders Orthoptera and Thysanoptera; and his
order Aptera was very different from ours.

These Orders may be briefly defined as follows,—the reader being asked

to recall the fact that by a mandibulate mouth we understand one in which
the mandibles, or the maxillæ, or both, are fitted for biting, crushing, or
grasping food; while the term suctorial implies that some of the mouth
parts are of a tubular form or are protrusible as a proboscis, which
assists, or protects, a more minute and delicate sucking apparatus:—

1. Aptera (ἀ without, πτερόν a wing). Wingless[108] Insects; mouth mandibulate or very

imperfectly suctorial. Metamorphosis very little.

2. Orthoptera (ὀρθός straight, πτερόν a wing). Four wings are present, the front pair
being coriaceous (leather-like), usually smaller than the other pair, which are of more
delicate texture, and contract in repose after the manner of a fan. Mouth mandibulate.
Metamorphosis slight.
3. Neuroptera (νεῦρον nerve, πτερόν a wing). Four wings of membranous consistency,
frequently with much network; the front pair not much, if at all, harder than the other pair,
the latter with but little or no fanlike action in closing. Mouth mandibulate. Metamorphosis
variable, but rarely slight.

4. Hymenoptera (ὑμήν membrane, πτερόν a wing). Four wings of membranous

consistency; the front pair larger than the hind, which are always small and do not fold
up in repose. Mouth mandibulate, sometimes provided also with a tubular proboscis.
Metamorphosis very great.

5. Coleoptera (κολεός sheath, πτερόν a wing). Four wings; the upper pair shell-like in
consistency, and forming cases which meet together over the back in an accurate line of
union, so as to entirely lose a winglike appearance, and to conceal the delicate
membranous hind pair. Mouth mandibulate. Metamorphosis great.

6. Lepidoptera (λεπίς scale, πτερόν a wing). Four large wings covered with scales.
Mouth suctorial. Metamorphosis great.

7. Diptera (δίς double, πτερόν a wing). Two membranous wings. Mouth suctorial, but
varying greatly. Metamorphosis very great.

8. Thysanoptera (θύσανος fringe, πτερόν a wing). Four very narrow fringed wings. Mouth
imperfectly suctorial. Metamorphosis slight.

9. Hemiptera (ἡμι half, πτερόν a wing). Four wings; the front pair either leather-like with
more membranous apex, or entirely parchment-like or membranous. Mouth perfectly
suctorial. Metamorphosis usually slight.

We must again ask the reader to bear in mind that numerous exceptions
exist to these characters in most of the great Orders; for instance,
wingless forms are not by any means rare in several of the Orders.

Before remarking further on this system we will briefly sketch two other
arrangements of the Orders of Insects, for which we are indebted to
Packard and Brauer.

Packard's Classification.

Packard has devoted much attention to the subject, and has published
two or three successive schemes, of which the following is the most